Using food network unfolding to evaluate food-web complexity in terms of biodiversity: theory and applications.

Food-web complexity often hinders disentangling functionally relevant aspects of food-web structure and its relationships to biodiversity. Here, we present a theoretical framework to evaluate food-web complexity in terms of biodiversity. Food network unfolding is a theoretical method to transform a complex food web into a linear food chain based on ecosystem processes. Based on this method, we can define three biodiversity indices, horizontal diversity (DH ), vertical diversity (DV ) and range diversity (DR ), which are associated with the species diversity within each trophic level, diversity of trophic levels, and diversity in resource use, respectively. These indices are related to Shannon's diversity index (H'), where H' = DH  + DV  - DR . Application of the framework to three riverine macroinvertebrate communities revealed that D indices, calculated from biomass and stable isotope features, captured well the anthropogenic, seasonal, or other within-site changes in food-web structures that could not be captured with H' alone.

Changes in the anatomy, morphology and mycorrhizal infection of fine root systems of Cryptomeria japonica in relation to stand ageing.

Biomass allocation to fine roots often increases under soil nutrient deficiency, but the fine root biomass does not often increase in old stands, even under nutrient limitation. Therefore, in old stands, the morphology, anatomy, branching architecture and mycorrhization of fine roots may compensate efficiently for nutrient acquisition by the low fine root biomass. In this study, changes in the morphology, anatomy and arbuscular mycorrhizal infection at each branching position of fine root clusters were evaluated in relation to stand age. A chronosequence (6­90 years of age) of stands in a Cryptomeria japonica D. Don plantation was used for these analyses. The fine root size parameters, such as length, weight and tip numbers of fine root clusters, increased with stand age. The specific root tip length (SRTL) decreased with increasing stand age, suggesting that the allocation to root active portions decreased with stand age. From the anatomical observation, the ephemeral root tips increased with stand age, suggesting that root tip turnover within a root cluster was high in old stands. The proportions of proto-xylem groups among branching positions indicated that the life cycles in branching hierarchy should be clearer in old stands than that in younger stands. The increasing in the mycorrhizal infection of root tips in old stands should enhance the root tip absorptive functions. The SRTL was correlated with the wood/needle ratio, suggesting that carbon limitation as the stand ages may result in decline of carbon allocation to maintain active root tips. However, increasing of the ephemeral tips and mycorrhizal infection rates may compensate the declines of tip allocation in old stands.

Terrestrial-aquatic linkage in stream food webs along a forest chronosequence: multi-isotopic evidence.

Long-term monitoring of ecosystem succession provides baseline data for conservation and management, as well as for understanding the dynamics of underlying biogeochemical processes. We examined the effects of deforestation and subsequent afforestation of a riparian forest of Japanese cedar (Cryptomeria japonica) on stable isotope ratios of carbon (δ¹³C) and nitrogen (δ¹5N) and natural abundances of radiocarbon (Δ¹4C) in stream biota in the Mt. Gomadan Experimental Forest and the Wakayama Forest Research Station, Kyoto University, central Japan. Macroinvertebrates, periphytic algae attached to rock surfaces (periphyton), and leaf litter of terrestrial plants were collected from six headwater streams with similar climate, topography, and bedrock geology, except for the stand ages of riparian forests (from 3 to 49 yr old in five stands and > 90 yr old in one reference stand). Light intensity and δ¹³C values of both periphyton and macroinvertebrates decreased synchronously with forest age in winter. A Bayesian mixing model indicates that periphyton contributions to the stream food webs are maximized in 23-yr-old forests. Except for grazers, most macroinvertebrates showed Δ¹4C values similar to those of terrestrial leaf litter, reflecting the influence of modern atmospheric CO2 Δ¹4C values. On the other hand, the Δ¹4C values of both periphyton and grazers (i.e., aquatic primary consumers) were significantly lower than that of modern atmospheric CO2, and were lowest in 23-yr-old forest stands. Previous studies show that root biomass of C. japonica peaks at 15-30 yr after planting. These evidences suggest that soil CO2 released by root respiration and dispersed by groundwater weathers carbonate substrata, and that dissolved inorganic carbon (DIC) with low Δ¹4C is incorporated into stream periphyton and some macroinvertebrates. The ecological response in the studied streams to clear-cutting and replanting of Japanese cedar is much slower (~20 yr) than the chemical response (< 5 yr). More than 50 yr is required for the food web structure to completely recover from clear-cutting. The ecological delay is attributed to several biogeochemical factors, the understanding of which is critical to integrated management of forest-stream continuum and the prediction of ecosystem resilience in response to environmental change.

Estimation of radioactive 137-cesium transportation by litterfall, stemflow and throughfall in the forests of Fukushima.

Since the Fukushima Dai-ichi Nuclear Power Plant accident in March 2011, large areas of the forests around Fukushima have become highly contaminated by radioactive nuclides. To predict the future dynamics of radioactive cesium ((137)Cs) in the forest catchment, it is important to measure each component of its movement within the forest. Two years after the accident, we estimated the annual transportation of (137)Cs from the forest canopy to the floor by litterfall, throughfall and stemflow. Seasonal variations in (137)Cs transportation and differences between forests types were also determined. The total amount of (137)Cs transported from the canopy to the floor in two deciduous and cedar plantation forests ranged between 3.9 and 11.0 kBq m(-2) year(-1). We also observed that (137)Cs transportation with litterfall increased in the defoliation period, simply because of the increased amount of litterfall. (137)Cs transportation with throughfall and stemflow increased in the rainy season, and (137)Cs flux by litterfall was higher in cedar plantation compared with that of mixed deciduous forest, while the opposite result was obtained for stemflow.

Consequence of altered nitrogen cycles in the coupled human and ecological system under changing climate: The need for long-term and site-based research.

Anthropogenically derived nitrogen (N) has a central role in global environmental changes, including climate change, biodiversity loss, air pollution, greenhouse gas emission, water pollution, as well as food production and human health. Current understanding of the biogeochemical processes that govern the N cycle in coupled human-ecological systems around the globe is drawn largely from the long-term ecological monitoring and experimental studies. Here, we review spatial and temporal patterns and trends in reactive N emissions, and the interactions between N and other important elements that dictate their delivery from terrestrial to aquatic ecosystems, and the impacts of N on biodiversity and human society. Integrated international and long-term collaborative studies covering research gaps will reduce uncertainties and promote further understanding of the nitrogen cycle in various ecosystems.

Nitrate isotopic composition reveals nitrogen deposition and transformation dynamics along the canopy-soil continuum of a suburban forest in Japan.

RATIONALE: Heavy nitrogen (N) deposition often causes high nitrate (NO3(-)) accumulation in soils in temperate forested ecosystems. To clarify the sources and production pathways of this NO3(-), we investigated NO3(-) isotope signatures in deposition processes along the canopy-soil continuum of a suburban forest in Japan. METHODS: The stable isotopes of N and oxygen (O) were used to trace the source and transformation dynamics of nitrate (NO3(-)) in two forest stands: a plantation of Cryptomeria japonica (coniferous tree; CJ) and a natural secondary forest of Quercus acutissima (broadleaf, deciduous tree; QA). The NO3(-) and ammonium (NH4(+)) concentrations were measured, as well as the δ(15)N and δ(18)O values of NO3(-), in rainfall, throughfall, stem flow, litter layer water, and soil water (10, 30, and 70 cm depths). RESULTS: Seasonal variations were observed in the δ(15)N values of throughfall and stem flow NO3(-) at both sites, and in the δ(18)O values of throughfall and stem flow NO3(-) at the QA site. The range in the δ(18)O values of rainfall and throughfall NO3(-) was large (65-70‰) but decreased dramatically to 2-5‰ in soil water at both sites. At the QA site, the δ(18)O values of stem flow NO3(-) decreased to 40‰ during several rain events, especially in the growing season. CONCLUSIONS: NO3(-) from atmospheric deposition was replaced by microbially generated NO3(-) mainly in the organic horizon and surface portion of the mineral soil under excess N deposition in this suburban forest. Microbial activity, including both immobilization and nitrification in organic-rich horizons near the surface, contributed to incorporating atmospheric NO3(-) quickly into the internal microbial N cycle. We also found evidence of microbial nitrification in the canopy of the QA stand during the growing season.

Differences between height- and light-dependent changes in shoot traits in five deciduous tree species.

The effects of tree height on shoot traits may in some cases differ in magnitude and direction from the effects of light. Nevertheless, general patterns of change in shoot traits in relation to variations in height and light have not so far been revealed. A comprehensive analysis of the differences between the effects of height and light on a range of leaf and shoot traits is important for the scaling of these traits to individual trees. We investigated the biomass allocation and structure of current-year shoots at the top of the crowns of five deciduous tree species in Japan. Height effect was investigated by comparing shoot traits among trees of different heights growing under a high light environment. The effects of light were examined by comparing saplings growing in high- and low-light environments. The effects of light were significant for most traits, while those of height were not significant for some traits. The magnitudes of the effects of light were larger than those of height for most traits related to biomass allocation. There was an extreme difference between the effects of height and light in the direction of change in the length of current-year shoots and in the number of standing leaves. The measures of both parameters increased with the increase in light, but decreased with the increase in tree height. Thus, the effects of height and light on diverse traits at the level of current-year shoots were not always similar. These results suggest that great care must be taken when scaling shoot traits from small trees to tall trees because the effects of height and light can be complex.

Nematomorph parasites indirectly alter the food web and ecosystem function of streams through behavioural manipulation of their cricket hosts.

Nematomorph parasites manipulate crickets to enter streams where the parasites reproduce. These manipulated crickets become a substantial food subsidy for stream fishes. We used a field experiment to investigate how this subsidy affects the stream community and ecosystem function. When crickets were available, predatory fish ate fewer benthic invertebrates. The resulting release of the benthic invertebrate community from fish predation indirectly decreased the biomass of benthic algae and slightly increased leaf break-down rate. This is the first experimental demonstration that host manipulation by a parasite can reorganise a community and alter ecosystem function. Nematomorphs are common, and many other parasites have dramatic effects on host phenotypes, suggesting that similar effects of parasites on ecosystems might be widespread.

Foliage nitrogen turnover: differences among nitrogen absorbed at different times by Quercus serrata saplings.

BACKGROUND AND AIMS: Nitrogen turnover within plants has been intensively studied to better understand nitrogen use strategies. However, differences among the nitrogen absorbed at different times are not completely understood and the fate of nitrogen absorbed during winter is largely uncharacterized. In the present study, nitrogen absorbed at different times of the year (growing season, winter and previous growing season) was traced, and the within-leaf nitrogen turnover of a temperate deciduous oak Quercus serrata was investigated. METHODS: The contributions of nitrogen absorbed at the three different times to leaf construction, translocation during the growing season, and the leaf-level resorption efficiency during leaf senescence were compared using (15)N. KEY RESULTS: Winter- and previous growing season-absorbed nitrogen significantly contributed to leaf construction, although the contribution was smaller than that of growing season-absorbed nitrogen. On the other hand, the leaf-level resorption efficiency of winter- and previous growing season-absorbed nitrogen was higher than that of growing season-absorbed nitrogen, suggesting that older nitrogen is better retained in leaves than recently absorbed nitrogen. CONCLUSIONS: The results demonstrate that nitrogen turnover in leaves varies with nitrogen absorption times. These findings are important for understanding plant nitrogen use strategies and nitrogen cycles in forest ecosystems.

Allocation of nitrogen within the crown during leaf expansion in Quercus serrata saplings.

Early season leaf growth requires a large amount of nitrogen, and the amount of N provided for new leaf development has been well tested. Although shoot position within the crown strongly influences leaf properties, little is known about absorbed and remobilized nitrogen allocation in the tree crown. Thus, we investigated differences in the allocation of recently absorbed nitrogen in the tree crown. To quantify nitrogen allocation, we conducted 15N tracer experiments using potted saplings of the temperate deciduous oak (Quercus serrata Thunb. ex. Murray). Allocation of 15N within the crown varied significantly: the top leaves received more remobilized nitrogen than did the lateral leaves, suggesting that remobilized nitrogen is predominantly allocated to the top shoots, which are important for height growth. On the other hand, the proportion of currently-absorbed nitrogen to total nitrogen in the lateral leaves was more than twice that in the top leaves. We also detected the input and the output of nitrogen in the top leaves after the completion of leaf expansion, indicating that significant nitrogen cycling occurs even after full leaf expansion.

Shoot development and extension of Quercus serrata saplings in response to insect damage and nutrient conditions.

BACKGROUND AND AIMS: Plants have the ability to compensate for damage caused by herbivores. This is important to plant growth, because a plant cannot always avoid damage, even if it has developed defence mechanisms against herbivores. In previous work, we elucidated the herbivory-induced compensatory response of Quercus (at both the individual shoot and whole sapling levels) in both low- and high-nutrient conditions throughout one growing season. In this study, we determine how the compensatory growth of Quercus serrata saplings is achieved at different nutrient levels. METHODS: Quercus serrata saplings were grown under controlled conditions. Length, number of leaves and percentage of leaf area lost on all extension units (EUs) were measured. KEY RESULTS: Both the probability of flushing and the length of subsequent EUs significantly increased with an increase in the length of the parent EU. The probability of flushing increased with an increase in leaf damage of the parent EU, but the length of subsequent EUs decreased. This indicates that EU growth is fundamentally regulated at the individual EU level. The probabilities of a second and third flush were significantly higher in plants in high-nutrient soil than those in low-nutrient soil. The subsequent EUs of damaged saplings were also significantly longer at high-nutrient conditions. CONCLUSIONS: An increase in the probability of flushes in response to herbivore damage is important for damaged saplings to produce new EUs; further, shortening the length of EUs helps to effectively reproduce foliage lost by herbivory. The probability of flushing also varied according to soil nutrient levels, suggesting that the compensatory growth of individual EUs in response to local damage levels is affected by the nutrients available to the whole sapling.

Soil functional responses to excess nitrogen inputs at global scale.

There is little evidence that nitrogen (N) cycling in the highly weathered, low-phosphorus (P), acidic soils found in Southern Hemisphere continents will differ greatly from that in North America and Europe. Evidence from the 'south' shows: the similarity in forms and temporal patterns in losses of N from different land uses; that the C:N ratios of the forest floor/litter layer from different continents are strongly predictive of a range of processes on a global scale; that generalizations based on Northern Hemisphere experience of the impact of N additions to 'P-limited' ecosystems are likely to fail for southern ecosystems where anatomical and physiological adaptation of native plants to low-P soils makes questionable the concept of 'P-limitation'; that the greatest threats in the 'south' are probably changes in land use that may greatly increase N inputs and turnover; that localized increases in N inputs produce similar effects to those seen in the 'north'.

Effects of NO3- availability on NO3- use in seedlings of three woody shrub species.

We determined the effects of short-term cultivation with various amounts of available nitrate nitrogen (NO3-) on NO3- use by woody shrub species. Nitrate concentration ([NO3-]) and nitrate reductase activity (NRA) were measured in leaves and roots of seedlings of Hydrangea hirta (Thunb.) Siebold, Lindera triloba (Sieb. et Zucc.) Blume and Pieris japonica (Thunb.) D. Don. Root [NO3-] increased with increasing NO3- supply in all species, whereas leaf [NO3-] remained low. There were significant correlations between [NO3-] in roots and leaves in all species, but no correlation was found between root NRA and leaf NRA. The low proportion of leaf NO3- assimilation to total NO3- assimilation in all species can be ascribed to the lack of NO3- transport from roots to leaves. In all species, root NRA increased with increasing NO3- supply until reaching a plateau. Species ranking based on maximum root NRA was H. hirta > L. triloba > P. japonica. Root NRA in P. japonica was low, even though root [NO3-] increased with NO3- supply, indicating that NO3- was not an effective N source for this species. The ranking also suggested that H. hirta depended more on NO3- as an N source than L. triloba. The increase in root NRA with increasing NO3- supply was greater in H. hirta than in L. triloba, possibly indicating that a change in NO3- availability has a stronger influence on NO3- use in H. hirta than in L. triloba.